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Meeting Society’s Needs for Education and Discovery:
A Survey of Eight Field Stations and Marine Laboratories in
the Southeastern United States
J. Christopher Havran1,*, Kirk A. Stowe2, Tom A. Blanchard3, Karen L. Kandl4,
Matthew E. Kimball5, Stephen C. Richter6, Hilary M. Swain7, Fred E. Lohrer7,
Dustin D. Angell7, and Theron M. Terhune II8
Abstract - Field stations and marine laboratories (FSMLs) are essential institutions for
natural history education and research. Recently, the National Research Council suggested
goals for FSMLs and that a metric be administered for their evaluation. We surveyed a
non-random cross-section of 8 southeastern US FSMLs in different ecosystems to evaluate
how they provide educational opportunities in accordance with National Research Council
recommendations pertaining to convergence, STEM education, and interdisciplinary education.
Survey responses were provided as narrative responses and through the completion of
a rubric. FSML representatives reported generally high mean scores in categories addressed
in the rubric. We observed the greatest variation in responses regarding the number of
interdisciplinary programs offered across FSMLs. Rubric responses associated with convergence
across programs, station culture, and facilities were relatively uniform. Although
the FSMLs surveyed exhibited variations in ecosystems serviced and programs offered, all
of the institutions provide a collaborative environment for individuals from multiple ages
and backgrounds.
Introduction
Field stations and marine laboratories (FSMLs) provide services to support
research, conservation, and natural history education by facilitating access to
natural settings for researchers, educators, students, and the public in a variety of
disciplines (Arvey and Reimer 1966, Billick and Price 2011, Eisner 1982, Kwok
2013, Michener et al. 2009, Richter et al. 2010). Onsite lodging, dedicated research
space, and legacy natural history collections at FSMLs can aid short- and long-term
ecological research projects and field-based educational programs (Billick et al.
2013, Porzig et al. 2011). Students can access FSMLs as a component of original
research, field schools, or as field-trip locations (Brown et al. 2011). Field stations
may also be accessible to the general public for education and recreation.
1Department of Biological Sciences, Campbell University, Buies Creek, NC 27506. 2210
Meadows Circle, Wixom, MI 48393. 3Department of Biological Sciences, University of
Tennessee at Martin, Martin, TN 38238. 4Highlands Biological Station, Highlands, NC
28741. 5Baruch Marine Field Laboratory, University of South Carolina, Georgetown, SC
29442. 6Department of Biological Sciences and Division of Natural Areas, Eastern Kentucky
University, Richmond, KY 40475. 7Archbold Biological Station, Venus FL, 33960.
8Tall Timbers Research Station and Land Conservancy, Tallahassee, FL 23212. *Corresponding
author - havran@campbell.edu.
Manuscript Editor: Eric Nagy
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Like all academic and research institutions, FSMLs face a variety of challenges
to continue meeting their commitment to conservation, research, and natural
history education. To this end, the National Research Council (NRC) recently
drafted a guide for future development of FSMLs (NRC 2014) that included
several recommendations pertaining to education. Among these guidelines, we
considered 2 to be particularly important: “Field station leaders should identify
and support the development of scientific and educational assets that harness their
stations’ unique qualities to address local, regional, national, and global challenges
by bringing together scientists from a number of disciplines, including the
social sciences, through what is now called convergence”; and “Universities and
other host institutions should expand opportunities at field stations to conduct
independent and collaborative research and active learning activities to increase
interest and persistence in STEM fields” (NRC 2014). In addition, the NRC also
recommended the establishment and implementation of metrics across regional
and national scales to assess the impact of field stations across mulitple factors,
including education (NRC 2014).
Numerous FSMLs that offer access to a variety of terrestrial and marine ecosystems
are located in the southeastern US (Arvey and Reimer 1966, Michener et
al. 2009), a region recently identified as the world’s 36th global diversity hotspot
(Noss et al. 2015). The region’s wide diversity of habitats has served to inspire and
cultivate natural history research for over a century (Odum 2002). Although there is
some coordination among FSMLs to set common goals through professional societies
(e.g., Organization of Biological Field Stations, National Association of Marine
Laboratories), only recently has there been an initiative to evaluate the function and
maximize the educational and research value of FSMLs at regional and national
scales (NRC 2014). During its 2014 meeting, the Education Committee of the Association
of Southeastern Biologists (ASB) sponsored a symposium to highlight
educational opportunities at FSMLs in the southeastern US. The symposium organizers
chose FSMLs that represented ecologically distinct areas dispersed across
the southeastern US (Havran and Stowe 2014). Representatives from 8 FSMLs in
the region shared information concerning educational activities occurring at their
respective institutions (Fig. 1).
In this paper, we evaluate how each FSML included in the ASB symposium currently
addresses the recommendations put forward by the NRC (2014). Although
these 8 FSMLs represented a non-random sample of institutions of different sizes,
ecosystems, and educational programs offered throughout the region, they represent
a cross-section of the diversity that exists. We developed a rubric for assessing
FSMLs’ facilities, educational opportunities, and NRC recommendations (Table 1).
The categories in the rubric focus both on the goals suggested by the NRC (2014)
pertaining to convergence and the facilities available for research and education. In
this paper we provide a mechanism for the assessment of education-related FSML
attributes and offer examples of how well existing FSMLs are meeting the goals
recommended by the NRC.
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Methods
Representatives from 8 southeastern FSMLs assessed their own institutions
qualitatively through a narrative response and quantitatively through the completion
of a rubric we developed. The numeric structure of the rubric should facilitate
its application across a variety of FSMLs. We acknowledge that the application of a
numeric rubric for use in assessing multifaceted, and often overlapping, attributes
of FSMLs may be problematic. We hope the narrative responses will serve as a
companion for interpreting the rubric scores.
The narrative responses are a summary of the oral presentations provided at the
2014 ASB symposium. Through narrative responses, FSML representatives identified
unique aspects of their respective FSML that promote (1) convergence and
(2) attention to science, technology, engineering, and mathematics (STEM) fields—
both components of the 2 NRC recommendations mentioned earlier. In their narratives,
FSML representatives (faculty and/or administrators) briefly summarized the
ecosystem serviced and how their FSML enhanced a variety of disciplines, increased
Figure 1. Distribution of FSMLs participating in the current survey (southeastern states
are shaded): ABS, Archbold Biological Station; BMFL, Baruch Marine Field Laboratory;
HBS, Highlands Biological Station; LCW, Lilley Cornett Woods Appalachian Ecological
Research Station; MEEL, Maywoods Environmental and Educational Laboratory; RLEFS,
Reelfoot Lake Environmental Field Station; TFEA, Taylor Fork Ecological Area; and
TTRS, Tall Timbers Research Station and Land Conservancy.
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interest and persistence in STEM fields by students, aided researchers and resource
managers, and fostered interest in natural history among members of the public.
The rubric we developed is similar to those used to assess institutions of higher
education (Timmerman et al. 2011), and we sought to determine if FSMLs were
adhering to the recommendations suggested by the NRC. The rubric (Table 1) assessed
different attributes pertaining to convergence, station culture, and facilities.
We employed the definitions of convergence provided by the NRC (2014). We
assessed convergence through 2 rubric responses: the “convergence: interdisciplinary
investigation” category encapsulated how well a particular FSML promotes
investigation across STEM disciplines, including life and health sciences, physical,
mathematical, and computational sciences, and/or engineering disciplines, whereas
the “convergence: programs” category addresses people, organizational structure,
culture, and research ecosystems associated with the FSML. A thriving station
culture is defined as “people of all ages come[ing] together at field stations” (NRC
2014). The “facilities” category includes teaching and research classrooms, laboratories,
and housing available for education.
Table 1. Rubric used for each of 4 categories to survey FSML representatives.
Numerical
value Level of performance
Convergence: Interdisciplinary investigation—degree to which institution promotes investigation
across disciplinary lines including life and health sciences, physical, mathematical, and computational
sciences, and/or engineering disciplines
0 Not successful: Does not promote interdisciplinary investigation
1 Moderately successful: Promotes investigation across 2 mentioned disciplines
2 Successful: Promotes investigation among 3 mentioned disciplines
3 Excels: Promotes investigation among more than 3 mentioned disciplines
Convergence: Programs—degree to which institution includes people, organizational structure, culture,
and research ecosystems
0 Not successful: Includes researchers working independently
1 Moderately successful: Includes researchers in an organizational (directed) structure
2 Successful: Includes researchers in an organizational structure with a view of the impact
of their research to non-discipline–specific issues
3 Excels: Includes all of previously mentioned facets in more than one system
Station culture—degree to which a station’s culture is the result of people of all ages and backgrounds
(K–12, undergraduate, graduate, senior scientists, public) coming together in a collaborative
environment
0 Not successful: Incorporates only a single group
1 Moderately successful: Incorporates 2 groups
2 Successful: Incorporates 3 or more groups
3 Excels: Incorporates 2 or more groups working collaboratively
Facilities—kinds of facilities available for education and research including classroom, laboratory,
and housing facilities
0 No facilities available
1 Classroom or laboratory facilities available
2 Classroom and laboratory facilities available
3 Classroom, laboratory, and housing facilities available
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Results
Narrative Responses of FSMLs
Archbold Biological Station (ABS). Education activities at Archbold Biological
Station draw from ABS research conducted throughout the regional watershed
known as the headwaters of the Everglades. ABS manages more than 7500 ha
of remarkable ecosystems in this strategic location, and provides nearly 5000 m2 of
laboratory, classroom, lodging, and public facilities that enable education activities
that draw from multidisciplinary research priorities and allow students to address
regional, national, and global challenges. With 1 education coordinator, 2 education
interns, and many volunteers, ABS offers innovative 3rd- through 5th-grade
curricula, summer camps, and high-school fellowships, which provide stimulating,
hands-on learning experiences for about 2000 schoolchildren annually. This
STEM-related education programming reaches an underserved rural community
that is 28% Latino and 19% African American. For decades, universities have
taught their undergraduate/graduate field courses at ABS (about 500 student days/
year). Post-baccalaureate research interns receive training in plant ecology, avian
ecology, entomology, agro-ecology, herpetology, and restoration ecology, conduct
an independent research project, and connect the relevance of their work to realworld
land-management and conservation issues.
ABS’s LEED® Platinum Learning Center, designed to inspire sustainable
living, opened in 2012. It welcomes the public with more than 100 signs and
displays, 6 self-guiding trails, tours, and public talks. The facility also hosts more
than 500 professional visitors annually for workshops, conservation meetings,
and scientific conferences. To expand its reach, ABS presents displays at Florida
festivals, participates in citizen science, engages with artists and writers, and has
a corps of volunteers.
Baruch Marine Field Laboratory (BMFL). The Baruch Marine Field Laboratory
is near a wide range of estuarine and coastal ecosystems that are representative of
the southeastern US. BMFL offers over 1950 m2 of classrooms and laboratories (indoor,
outdoor, and seawater), as well as conference facilities and overnight housing
for ~70 individuals. Major research themes at BMFL have focused on topics such
as water quality/chemistry, marsh-grass growth and production, sediment dynamics
and geomorphology, and the ecology of fish and shellfish.
Education activities at BMFL are centered on undergraduate- and graduate-student
involvement in field trips, onsite courses, and participation in ongoing research
through internships. More than 710 projects have been completed in the last 45 years,
many involving students from the physical, chemical, and life sciences. Multi-week
summer field-courses are regularly offered onsite through the University of South
Carolina Marine Science Program, providing immersive experiential learning for as
many as 20 upper-level undergraduates per course. In addition to field trips for undergraduate
and graduate classes from disciplines as varied as art, law, and biology,
BMFL facilities are regularly used for non-traditional education activities, including
teacher workshops, training programs for coastal-zone managers, and outreach activities
for children and adults. These activities are designed to provide discovery-based
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learning opportunities, generate and increase interests in marine science and other
STEM fields, and convey the results of scientific research conducted at BMFL and
elsewhere to the community and K–12 classrooms.
Eastern Kentucky University (EKU) field stations. Eastern Kentucky University
owns and operates 3 field stations: Taylor Fork Ecological Area (TFEA),
Maywoods Environmental and Educational Laboratory (MEEL), and Lilley Cornett
Woods Appalachian Ecological Research Station (LCW). Eastern Kentucky
University stations are members of a newly formed collaborative network, the
Kentucky Organization of Field Stations (KOFS). The EKU Division of Natural
Areas manages these stations and facilities, makes them available for research, and
provides funds to support students, environmental monitoring, research-informed
training, interdisciplinary collegiate education, K–12 environmental education, and
community outreach. The EKU field stations are guided by an integrative mission
that primarily focuses on multidisciplinary research, teaching, and outreach.
Taylor Fork Ecological Area is adjacent to EKU’s campus and is comprised
of 24 ha of old-field habitat undergoing ecological succession and restoration.
Habitats present include small patches of trees, streams, and constructed wetlands.
Because of its proximity to campus, EKU faculty and students frequently use the
site for field trips, research projects, service-learning projects, biological monitoring,
and educational outreach.
Maywoods Environmental and Educational Laboratory is a 688-ha forested site
35 km from EKU. Habitats include oak–hickory forest scattered across a series of
knobs, restored tall-grass prairie, wetlands, and a 5-ha lake. The station has teaching
and lodging facilities and is used regularly for teaching, research, and outreach.
The Division of Natural Areas has hosted college field-courses at Maywoods in
the summer and has offered science programming there to local schools for over a
decade, averaging 1600 students per year for the past 4 years.
The LCW is a 223-ha forested site located 221 km from campus. The first oldgrowth
forest preserved in Kentucky occupies over half of the station’s land. Habitats
available for study at the station include mixed mesophytic forest with multiple community
types, wetlands, first-order and larger streams, and some open grassy areas
around developed sites. Facilities for teaching, research, and housing are available
onsite as well as on an adjacent 32-ha property within a secondary-growth forest. The
staff host researchers and college classes from EKU and other institutions, agencies,
local school groups, and provide guided tours for the general public.
Highlands Biological Station (HBS). The mission of Highlands Biological Station
is to foster research and education focused on the rich natural heritage of the
Highlands Plateau in North Carolina. HBS has promoted both research and education
for local residents, visitors, and academics since 1927. The station’s research legacy
lies in salamander biology, but HBS researchers also study the taxonomy, systematics,
ecology, evolution, and conservation of many groups of terrestrial and aquatic
organisms. Each summer, HBS offers accredited field-biology courses at advanced
undergraduate and graduate levels, as well as introductory workshops on topics ranging
from pure science to arts and science. HBS also offers workshops for the general
public that integrate biology and other fields of study, including art and literature.
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Outreach education at HBS is based in the HBS Nature Center, which has a
decades-long tradition of day and evening programs that provide science-based
enrichment for the community, regional schools, and tourists. The Nature Center
offers over 20 different day-programs that cover a variety of science and nature
classes and outreach programs for grades pre-K–12. Most of the classes are
science-based, but integrate other subject areas such as mathematics, geography,
and language arts. The station has also instituted “science a la carte”, a program in
which summer researchers spend a few hours of their time discussing their research
with visitors to HBS.
Reelfoot Lake Environmental Field Station (RLEFS). Reelfoot Lake Environmental
Field Station offers a variety of unique research, educational, and outreach
opportunities due to its proximity to terrestrial and aquatic habitats. Reelfoot Lake
is the largest of Tennessee’s natural lakes and includes ~5500 ha of open water and
associated aquatic habitats. The combination of the lake’s eutrophic nature, relative
youth, and tectonic origin provide an unusual set of physical and biological features.
The primary mission of RLEFS is to serve as a resource for undergraduate
education and research in the Reelfoot Lake area. Although graduate credit can
be obtained, all courses focus on undergraduates, and are specifically designed to
deliver experience with the practical tools and methods routinely used by professionals
in their respective fields. The station offers 1 summer course (Geology of
the Reelfoot Lake Area) in STEM-based subjects outside of the biological sciences.
The station director is on the advisory board of the University of Tennessee at
Martin’s (UTM) West Tennessee STEM Center for Learning and actively recruits
potential instructors, students, and researchers in STEM-based fields.
The station’s annual open-house is intended to stimulate public interest in natural
history science and to increase awareness of environmental issues relevant to
the long-term health of the Reelfoot Lake system. RLEFS and UTM partner with
the Discovery Park of America in Union City, TN, to sponsor an internship during
which a student assists with the care and demonstration of live plants and animals
common to the Reelfoot Lake area.
Tall Timbers Research Station and Land Conservancy (TTRS). Tall Timbers,
a stand-alone non-profit institution, is in Leon County, FL, and is situated in a
unique geophysical region commonly referred to as the “Red Hills”. Most of the
property consists of upland pine forest intermixed with hardwood hammocks and
annually disked fallow fields. TTRS is internationally regarded as an information
resource in the areas of fire ecology, game-bird management, vertebrate ecology,
and forestry. Station representatives update and help maintain the E.V. Komarek
Fire Ecology Database, an open-access directory for fire-ecology references, citations,
and documents.
TTRS regularly hosts land managers’ luncheons, field days, and field tours,
while staff teach several courses each year germane to prescribed fire, geospatial
techniques and analysis, and upland-game and land management. In addition,
TTRS is affiliated with more than 25 different academic institutions that provide
collaborative graduate and undergraduate educational and research opportunities
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in the classroom and in the field. The field station engages the public by surveying
communities, organizations, and individuals; public advocacy; lobbying; and the
hosting of public events. TTRS also directly communicates with the public through
the production of occasional television shows with Georgia Public Broadcasting,
videos, and magazine articles.
Rubric responses
Rubric responses of FSML representatives are presented in Table 2. The total
rubric scores for the FSMLs range from 6 at RLEFS to 12 at ABS, LCW, and TTRS,
demonstrating variation in how well FSML representatives think they are meeting
the NRC recommendations addressed in the rubric. Category means across FSMLs
range from 2.1–2.9 (maximum possible response = 3).
The mean for “convergence: interdisciplinary investigation” category was
the lowest of all categories (mean = 2.1). There was also quite a bit of variation in
scores in this category (modes = 2, 3). ABS, LCW, and TTRS reported performing
well in this category because they facilitate investigation across more than 3 of the
mentioned disciplines (Table 2). With the exception of RLEFS, most station representatives
reported high scores in the “convergence: programs” category.
For the “station culture” and “facilities” categories, most stations reported
the maximum score of 3. RLEFS reported a 2 for station culture, indicating that it
incorporates 3 or more groups of people with different backgrounds. Taylor Fork
Ecological Area reported a 0 in the “facilities” category, indicating that they do not
offer any classroom, laboratory, or housing facilities.
Discussion
The FSMLs included in this survey are located throughout all biogeographic
regions in the southeastern US as described by Sayre et al. (2009). The Central
Interior and Appalachian Biogeographic Regions are represented by TFEA, MEEL,
LCW, and HBS; RLEFS, BMFL, and TTRS are in the Gulf and Atlantic Coastal
Plain. ABS, in south-central Florida, is on the boundary of the Coastal Plain and the
Caribbean Biogeographic Region (Sayer et al. 2009). In addition to being distributed
across these 3 major biogeographic regions, the field stations provide access
to a variety of unique terrestrial, freshwater, and marine ecosystems within these
broader biogeographic regions.
Table 2. Responses of individual FSML representatives regarding how well their FSML meets NRC
recommendations (abbreviations follow those from Fig. 1).
Category
ABS BMFL TFEA MEEL LCW HBS RLEFS TTRS x̅ Mode(s)
Convergence: Interdisciplinary 3 2 1 2 3 2 1 3 2.1 2, 3
investigation
Convergence: Programs 3 3 3 3 3 3 0 3 2.6 3
Station culture 3 3 3 3 3 3 2 3 2.9 3
Facilities 3 3 0 3 3 3 3 3 2.6 3
Total 12 11 7 11 12 11 6 12 - -
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We assessed convergence through scores for 2 rubric categories: “convergence:
interdisciplinary investigation” and “convergence: programs”. The greatest amount
of variation seen in scores for any rubric category was in “convergence: interdisciplinary
investigations”. Though some of the surveyed FSMLs do not balance
their interdisciplinary investigations equally within and between science and nonscience
disciplines, their strength is that they promote at least some investigation
across the sciences and the arts.
With the exception of RLEFS, all station representatives reported the maximum
score in the “convergence: programs” category, indicating that they include researchers
in an organizational structure with a view of the impact of their research
to non-discipline–specific issues in more than 1 system. This finding does not
indicate that RLEFS is not contributing to broader research in the area, but that
researchers are working independently, not through a directed organizational structure
at the field station.
People of all ages and backgrounds (K–12, undergraduate, graduate, senior scientists,
general public) coming together in a collaborative environment contribute
to the “station culture” category (Table 2). As with the “convergence: programs”
category, the scores for the “station culture” category were all reported at the maximum
of 3 with the exception of RLEFS, whose FSML representative reported a 2
for “station culture”, indicating that it incorporated 3 or more groups of people of
different ages and backgrounds.
The score for “facilities” was high for all but 1 of the FSMLs surveyed. Only
TFEA reported a 0 in this category. In the narrative, the respondent indicated this
FSML is located adjacent to the campus of EKU. Some FSMLs may lack facilities
because they rely on their proximity to university or college campuses, which can
facilitate easy access to field and lab equipment. Despite reporting a 0 in the “facilities”
category, TFEA reported high scores in both the “convergence: programs” and
“station culture” categories. This result suggests that onsite classroom, housing,
and laboratory facilities are not necessarily important for bringing together different
groups of people in a collaborative environment.
Various activities are conducted to increase interest and persistence in STEM
fields. Many of the research topics explored by FSMLs are likely to significantly
overlap with STEM fields. Through public outreach and participation in K–12
educational programs, FSMLs are able to support STEM-related disciplines. It is
likely that many FSMLs will be located in relatively undeveloped and rural areas;
thus, the education provided at stations has great potential benefit for otherwise
underserved rural communities. Directors and staff may also help to guide regional
STEM initiatives by serving on STEM advisory councils, as was indicated in the
narrative response for RLEFS.
In providing access to a variety of ecosystems in the southeastern US, the surveyed
FSMLs indicated they service a variety of disciplines beyond STEM fields
including art, law, language arts, and geography. Some of these non-STEM
fields are incorporated in public outreach opportunities. Methods employed to
provide outreach and education to the general public and resource managers may
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involve informational displays in FSML welcome centers, workshops, researcher
interactions, and television broadcasts.
The generally high scores in our survey suggest that the surveyed southeastern
FSMLs are successfully meeting NRC recommendations. However, FSMLs
can always make further progess toward meeting NRC recommendations. The
“convergence: interdisciplinary investigation” category received the lowest mean
score among all categories; TFEA, scored a 1. Despite this low score, TFEA may
ultimately benefit from being part of the KOFS, which may be able to to share resources
across the state. We recommend that other states follow the model of the
KOFS to provide a regional network of FSMLs (Richter et al. 2010).
The metrics presented in this survey are the first attempt at evaluating southeastern
FSMLs in terms of the recommendations provided by the NRC (2014),
especially those involving convergence across disciplines. This concept is
incredibly important to natural history education (Schubel 2015) because it
provides the opportunity to attract an audience of diverse learners to FSMLs.
Use of an established set of evaluation metrics pertaining to FSMLs will provide
data needed to encourage their development, overcome potential challenges to
natural history education, and meet their full potential (Krupa 2000, Schubel
2015, Wilson 1982).
We caution that the metrics developed here are an initial attempt at evaluating
FSMLs using those of the southeastern US as a case study. The FSMLs included
were invited by the organizers of the ASB Education Committee symposium; as
such, they represent a potentially biased sample of FSMLs. The fact that faculty
and/or FSML administrators were tasked with assessing their own stations may
also have biased the results. However, we feel that the information provided by the
FSMLs will help to inform other stations and provide a reference for future assessments
of FSMLs.
We recommend that an independent, impartial observer conduct broad assessments
of FSMLs. Future assessments could also be limited to FSMLs in a given
ecosystem type (marine, mountain, freshwater, etc.), or within 1 biogeographic region
because each ecosystem may require a different suite of facilities that are not
comparable across all FSMLs in a geographic region. Additional surveys might also
attempt to determine how well a FSML meets the mission of its home institution, if
it has one. Furthermore, additional surveys should attempt to increase the number
of FSMLs included in the assessment.
The diversity of ecosystems of the southeastern US is reflected in the diversity
and missions of its FSMLs. Southeastern FSMLs provide educational
opportunities across a variety of disciplines in the arts and sciences. Their attention
to STEM education reinforces and inspires interest in natural history among
undergraduate, graduate, and pre-K–12 students. Despite differences in ecosystems
serviced and number of programs offered at FSMLs, the most important
commonality of these institutions is that they inspire interest in, and research
focused on, natural history.
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Acknowledgments
We thank the Executive Committee of the Association of Southeastern Biologists and
an anonymous donor for providing funding for the symposium that provided the basis for
this article. Zack Murrell provided invaluable encouragement during the drafting of early
manuscripts. Melinda Wilder and others provided helpful comments on this manuscript.
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